Treasures that scientists keep on ice

Honouring Alexander

Maybe it’s uncool to talk about heroes in science these days, because everyone is poised on others’ shoulders, but “Neill” (Robert McNeill) Alexander is undeniably a hero to many researchers in biomechanics and other strands of biology. Our lab probably wouldn’t exist without his pervasive influence- he has personally inspired many researchers to dive into biomechanics, and he has raised the profile of this field and championed its importance and principles like no other one individual. Often it feels like we’re just refining answers to questions he already answered. His influence extends not only to comparative biomechanics and not only around his UK home, but also –via his many, many books on biology, anatomy and related areas, in addition to his research, editorial work and public engagement with science– to much of the life sciences worldwide.

Sure, one could (and with great humility I’m sure Alexander would) mention others like Galileo and Marey and Muybridge and Fenn and Gray and Manter who came before him and did have a profound impact on the field. Alexander can, regardless, easily be mentioned in the same breath as luminaries of muscle physiology such as AV Hill and even Andrew + Julian Huxley. But I think many would agree that Alexander, despite coming later to the field, had a singular impact on this young field of comparative biomechanics. That impact began in the 1970s, when Dick Taylor and colleagues in comparative physiology were also exploding onto the scene with work at the Concord Field Station at Harvard University, and together biomechanics research there, in the UK, elsewhere in Europe and the world truly hit its stride, with momentum continuing today. I’m trying to think of some women who played a major role in the early history of biomechanics but it was characteristically a woefully male-dominated field. That balance has shifted from the 1970s to today, and my generation would cite luminaries such as Mimi Koehl as key influences. There are many female or non-white-male biomechanics researchers today that are stars in the field, so there seems to have been progress in diversifying this discipline’s population.

Hence, honouring Alexander’s impact on science, today our college gave Neill an honorary doctorate of science (DSc). Last year, I also helped organize a symposium at the Society for Vertebrate Paleontology’s conference in Berlin that honoured his impact specifically on palaeontology, too- compare his book “The Dynamics of Dinosaurs and Other Extinct Giants” to current work and you’ll see what fuelled much of that ongoing work, and how far/not far we’ve come since ~1989. Even 10 years later, his “Principles of Animal Locomotion“, with Biewener’s “Animal Locomotion“, remains one of the best books about our field (locomotion-wise; Vogel’s Comparative Biomechanics more broadly) , and his educational CD “How Animals Move“, if you can get it and make it work on your computer, is uniquely wonderful, with games and videos and tutorials that still would hold up well as compelling introductions to animal biomechanics. Indeed, I’ve counted at least 20 books penned by Alexander, including “Bones: The Unity of Form and Function” (under-appreciated, with gorgeous photos of skeletal morphology!).

1970s Alexander, with a sauropod leg.

And then there are the papers. I have no idea how many papers Neill has written –again and again I come across papers of his that I’ve never seen before. I tried to find out from the Leeds website how many papers he has, but they’re equally dumbfounded. I did manage to count 38 publications in Nature, starting in 1963 with “Frontal Foramina and Tripodes of the Characin Crenuchus,” and 6 in Science. So I think we can be safe in assuming that he has written everything that could be written in biomechanics, and we’re just playing catchup to his unique genius.

Seriously though, Alexander has some awesome publications stemming back over 50 years. I’m a big fan of his early work on land animals, such as with Calow in 1973 on “A mechanical analysis of a hind leg of a frog” and his paper “The mechanics of jumping by a dog” in 1974, which did groundbreaking integrations of quantitative anatomy and biomechanics. These papers kickstarted what today is the study of muscle architecture, which our lab (including my team) has published extensively on, for example. They also pioneered the integration of these anatomical data with simple theoretical models of locomotor mechanics, likewise enabling many researchers like me to ride on Alexander’s coattails. Indeed, while biomechanics often tends to veer into the abstract “assume a spherical horse”, away from anatomy and real organisms, Alexander managed to keep a focus on how anatomy and behaviour are related in whole animals, via biomechanics. As an anatomist as well as a biomechanist, I applaud that.

Alexander has researched areas as diverse as how fish swim, how dinosaurs ran, how elastic mechanisms make animal movement more efficient, how to model the form and function of animals (see his book “Optima for Animals” for optimization approaches he disseminated, typifying his elegant style of making complex maths seem simple and simple maths impressively powerful) and how animals walk and run, often as sole author. In these and other areas he has codified fundamental principles that help us understand how much in common many species have due to inescapable biomechanical constraints such as gravity, and how these principles can inspire robotic design or improvements in human/animal care such as prosthetics. Neill has also been a passionate science communicator, advising numerous documentaries on television.

~1990s Alexander, with model dinosaurs used to estimate mass and centre of mass.

Alexander’s “Dynamics of Dinosaurs” book, one of my favourites in my whole collection, is remarkably accessible in its communication of complex quantitative methods and data, which arguably has enhanced its impact on palaeontologists. Alexander’s other influences on palaeobiology include highly regarded reviews of jaw/feeding mechanics in fossil vertebrates (influencing the future application of finite element analysis to palaeontology), considerations of digestion and other aspects of metabolism, analysis of vertebral joint mechanics, and much more. Additionally, he conducted pioneering analyses of allometric (size-related) scaling patterns in extant (and extinct; e.g. the moa) animals that continue to be cited today as valuable datasets with influential conclusions, by a wide array of studies including palaeontology—arguably, he helped compel palaeontologists to contribute more new data on extant animals via studies like these.

Neill Alexander did his MSc and PhD at Cambridge, followed by a DSc at the University of Wales, a Lecturer post at Bangor University and finally settling at the University of Leeds in 1969, where he remained until his retirement in 1999, although he maintains a Visiting Professorship there. I had the great pleasure of visiting him at his home in Leeds in 2014; a memory I will treasure forever, as I had the chance to chat 1-on-1 with him for some hours. He has been Secretary of the Zoological Society of London throughout most of the 1990s, President of the Society for Experimental Biology and International Society of Vertebrate Morphologists, long championing the fertile association of biomechanics with zoology, evolutionary biology and anatomy. More recently, he was a main editor of Proceedings of the Royal Society B for six years.

Many people I’ve spoken to about Neill before have stories of how he asked a single simple question at their talk, poster or peer review stage of publication, and how much that excited them to have attracted his sincere interest in their research. They tend to also speak of how that question cut to the core of their research and gave them a facepalm moment where they thought “why didn’t I think of that?”, but how he also asked that question in a nice way that didn’t disembowel them. I think that those recalling such experiences with Neill would agree that he is a professorial Professor: a model of senior mentorship in terms of how he can advise colleagues in a supportive, constructive and warmly authoritative, scholarly way. For a fairly recent example of his uniquely introspective and concise, see the little treasure “Hopes and Fears for Biomechanics”, a ~2005 lecture you can find here. I really like the “Fears” part. I share those fears- and maybe embody them at times…

My visit with RMcNeill Alexander in 2014.

Perhaps I have gushed enough, but I could go on! Professor RMcNeill Alexander, to summarise the prodigious extent of his research, is to biomechanics as Darwin is to biology as a whole. One could make a strong case for him being one of the most influential modern biologists. He is recognised for this by his status as a Fellow of the Royal Society (since 1987), and a CBE award, among many other accolades, accreditations and awards. And, if you’ve met him, you know that he is a gentle, humble, naturally curious and enthusiastic chap who instils a feeling of awe nonetheless, and still loves to talk about science and keeps abreast of developments in the field. And as the RVC is honouring Neill today, it is timely for me to honour him in this blog post. There can never be another giant in biomechanics like Alexander, and we should be thankful for the broad scientific shoulders upon which we are now, as a field, poised.

I hope others will chime in with comments below to share their own stories.

Thank you for this, what a great post!
I’ve never met him, but he’s had an influence on me since I was about 12. The summer I turned 12 my dad got me a copy of Dynamics of Dinosaurs for my birthday. I fell in love with biomechanics and subsequently read most of his books in high school and college, along with the classics by Schmidt-Nielson and Vogel. I don’t know what I would have become without Optima for Animals or other likeminded books, and it’s those early exposures to it that made the difference.

Thanks, John, for this eloquent and richly deserved tribute to a true giant. The mark of Alexander is that he could do such far-reaching work (and so much of it) but also condense so much into so short and readable a book as “Dynamics of Dinosaurs and other Extinct Giants” for a general audience. How many academics could write a book on their specialist subject that my son could have read and enjoyed at the age of 10 or 13?

Like you, I consider that book to be one of the jewels of my collection. It’s probably no exaggeration to say that it’s one of the two books (along with The Dinosaur Heresies) that made me a palaeontologist.

In 2003 at the American Society of Biomechanics meeting in Toledo, Neill talked with my graduate students at the opening reception giving them advice about science. It was fantastic. Claire Farley had introduced me to him years before. I saw him standing in a corner at the reception by himself and took the opportunity to re-introduce myself to him and talk with him. I noticed a table of my graduate students were on the other side of the room, eyes fixed on Neill and me with their jaws on the floor. I asked Neill if he wouldn’t mind sitting with my students for a few minutes to chat with them. He stayed about about an hour and had some very valuable pieces of advice. One in particular: when you are starting up your own lab, find something nobody else is working on and make yourself the world’s expert. He has inspired a lot of biomechanists around the world, including ones that don’t work on comparative biomechanics.

I should add something serious too. It may be almost impossible for today’s students to realize it, but before Alexander (and Dawson & Taylor’s kangaroo paper), people thought of tendons as rigid structures with muscles undergoing large stretch-shorten cycles. Legs = springs : Mind = blown. Running, hopping, trotting etc. only make sense when we think of tendons as springs.

Dear colleagues!
Let me introduce myself. I was the editor and project leader of the “How Animals Move” CD.
In the late 1992, the friend of mine, who graduated together with me from the Moscow Lomonosov State University, and then worked in laboratory of Nikolay Kokshaysky (do you remember his pioneering visualization of the bird wake in 1979?)… so, the friend of mine called me by phone to ask if I want to work with Professor Alexander? I immediately answered that I definitely want, because by that time the article on kangaroo by Alexander and Vernon was my favourite one (as well as Romer’s “Locomotor apparatus of certain…”).
So, the project started, and I worked under the distant guidance of Neill for almost two years. Only one day we spent together in London, discussing the final efforts on the project, and then, only one time we communicated by phone. In spite of so brief communication, I regard Neill as my teacher.
If you are interested, I can tell more details of this fantastic project.

I am very sorry for enormous delay, but finally I’ve gathered my mind and tell the tale. Indeed, it is the tale, because only Nick Maris, the founder of Maris Multimedia Ltd. (later, Maris Technologies Ltd.) knows how it really was.

Prologue
Nick Maris is a British businessman and millionaire, whose family business was yacht-building for other millionaires. He is a very well educated person (Cambridge or Oxford, I don’t remember) and a poet and adventurer in his soul. He decided to found a new business, which was the development of educational multimedia CD-ROMs by cheap and talented Russian staff. It was in the very beginning of 1990-s – the time of maximum freedom and minimum money here in Russia. It is a mystery how, but Nick Maris fell from the British heaven on the Russian earth exactly in our Space Mission Control Center, the Korolev town near Moscow (Sergey Korolev is the name of the head of Russian cosmonautics), and exactly when many young physicians, astronomers and programmers were running away from the Space Center in search for better salary. Nick caught 5 or 7 of the most talented of them, added, somehow, two young Russian artists, and for funny money (which were not funny at all here at that time) they created, in a year, the famous first version of astronomic CD-ROM “Readshift” http://archaic.maris.com/content/indexe5cc.html . The title implies both the Doppler effect and the Russian origin of the staff and placement of the studio.
When development of the Readshift project was successfully coming to its end, Nick began to think on the topics for future multimedia discs. He was fascinated by the fresh-published in 1992 “Exploring Biomechanics: Animals in Motion” by Neill Alexander (don’t you remember its super cover with basilisk running on water?), came to him, suggested to be the author of a multimedia CD-ROM based on this book and asked him, who of Russian biologists can be a project leader for production of the CD-ROM in his Russian studio. Neill named Nikolay Kokshaysky. In fact, among Russian biomechanicians of animals, only Kokshaysky was well known on the other side of the Iron Curtain due to his pioneering article in Nature on visualization of the wake structure of a flying bird (Jeremy Rayner published similar results but 5 years later).

Action
By the end of 1992, Maris found Kokshaysky in Moscow, but he refused to participate in the project because of his age (he was born in 1931 and died in 2000), and suggested my person for the project leader, instead. I’ve found 2 programmers and 3 artists and we did the job in 18 months.
I am very happy to have met both Mr. Maris and Prof. Alexander. In the Soviet Union, due to persistent care of the government apparatus, most people including myself were staying in some underdeveloped condition – call it wild or child. Work with Nick Maris made me a self-contained person. Work with Neill Alexander and one brief meeting with him… well, it was a pivot. What impressed me most of all, were Neill’s manuscripts for our CD. They were hand-written on A4 pages with clear tall script and almost WITHOUT ANY STRIKEOUTS OR OTHER CORRECTIONS over hundreds of pages! What a clear mind he has! Also, Neill helped me to publish my first article in English (on the energetic profit of the three-segment limb), in the Journal of Theoretical Biology. I remember that he wrote in his review, that my article “strikes me (him) as a valuable contribution to our understanding of legged locomotion”. I was truly happy!

Epilogue
I still use our “How Animals Move” CD-ROM to teach my students. Under Windows 7 or later it is necessary to install Windows Virtual PC and then Windows XP mode, put the CD in the drive and run it.

Great story, thank you so much Alex! It is well worth the minor delay. That fills in a big gap in my understanding of how that amazing CD-ROM came about. And yes I do remember that book Exploring Biomechanics; I still know exactly where it is on my bookshelf. 🙂 But there still is no better teaching tool in comparative biomechanics than that CD-ROM!

[…] (inverse dynamics analysis of musculoskeletal mechanics), and owed a lot to simpler approaches by RMcNeill Alexander and others, but probably was published (and gained me some notoriety/infamy) because it answered a […]

Dear John! I’ve downloaded Neill’s “Hopes and fears” lecture from your site, and I’m going to present it, as it is, to my students as a final lecture in a training course on Biomechanics, which I invent in the Moscow State University for the first time this Spring. But I don’t understand exactly what did Neill say for each slide of his presentation. Don’t you have some record of his lecture – video, audio, or text? I would be greatly indebted indeed.

Yesterday I’ve shown Neill’s presentation on “Hopes and Fears” as the last lecture in my course on biomechanics.
It seems to have gone fine. I’ve managed to explain every slide.
Most of the time I devoted to explanation of Kram & Taylor hypothesis and the first slide plotting
metabolic cost of transport in running together with mechanical energy of the center of mass (taken from Full & Tu, 1991).
The question which Neill definitely meant here is why the mass-specific cost of running falls as mass^(-1/3),
while mechanical energy fluctuations per meter per kilogram remain almost constant.
I am sure that Neill was greatly suspicious about Kram & Taylor hypothesis, because,
on the one hand, it is the only hypothesis that explains the sacral law of mass^(-1/3) decrease of the metabolic cost,
and on the other hand, it is based on a strange assumtion that the cost of muscular force overrides the cost of muscular work.

Thinking after the lecture on the problem, I’ve, seems to me, guessed the sacral puzzle of mass^(-1/3).
Just look.
What is the unit of mass-specific metabolic cost of transport?
Yes, it is Joule/(meter*kg) = Newton*meter/(meter*kg) = (kg*meter/second^2)/kg = meter/second^2
This is the unit of acceleration.
So, on the graph from Full & Tu, 1991 it is more adequate to place, instead of the plot of mechanical energy of the center of mass,
the plot of typical accelerations in running.
It is very easy to imagine the line of acceleration on this graph.
Indeed, acceleration = force/mass = (work/shift)/mass
Let it be the average vertical (antigravitational) acceleration of the center of mass in the contact phase.
The mass is the mass of the body, the force is the average vertical ground reaction force,
shift is the distance over which the force is applied and it is definitely proportional to leg length,
while work is proportional to muscular mass involved in antigravitational action.
Thus, acceleration ~ (muscular mass)/(leg length/body mass).
We can easily assume that in all running animals musculature is proportionately involved in mechanical work against gravity.
So, acceleration ~ 1/(leg length) ~ mass^(-1/3) ~ (mass-specific metabolic cost of transport)

Taking into account the same units of acceleration and cost,
it seems appropriate to estimate the efficiency by the ratio: (acceleration/mass-specific metabolic cost of transport)
Both of them fall as mass^(-1/3), and so the muscular efficiency is size-independent in fact.

What do you think about it? Would Neill be happy of this simple solution?

There were two great beards at Leeds when I was an undergraduate in the mid 1970s – McNeill Alexander, Professor of Zoology and Harold Woolhouse, Professor of Botany. There was a weekly seminar series with external speakers that alternated between a botanist and a zoologist. As undergraduates we attended these religiously just so we could see these two great men put the visitors on the spot – McNeill Alexander always used to ask the first question after the botanist visitor had finished, prefacing his question with the words ‘Well, I’m only a zoologist, but..” and would the proceed to ask an extremely perceptive and challenging question. At the end of each zoological seminar, Woolhouse would hoist himself to his feet and say “Well I’m only a botanist, but…”

I am so very sorry to hear this. He was my Prof when I was an undergraduate in the early 70s and I undertook a project with him on Mole digging mechanics and he was wonderfully tolerant. I think he saw potential in people and when I wanted to stay in Leeds to do a PhD he told me to go elsewhere. Wise advice that I have passed on. He never showed any favouritism and I had several “discussions” about science with him and interpretation of data. Always a gentleman, always a scientist and such a clear thinker.

For those that don’t know, Professor Emeritus Robert McNeill (“Neill”) Alexander passed away this week after a short illness. It is terribly sad news and he will be greatly missed by a vast number of his colleagues and former students, whom we’ve already heard many wonderful stories from about how he was a shining light of kindness and scholarship in their lives. I encourage anyone wanting to share their stories about interactions with Neill to add them here in the Comments, so that we can continue to honour him.